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I UNCOVERING the CONSEQUENCES of CO UNCOVERING THE CONSEQUENCES OF CO-FLOWERING AND POLLINATOR SHARING: EFFECTS OF LOCAL COMMUNITY CONTEXT ON POLLEN TRANSFER DYNAMICS, FEMALE REPRODUCTIVE SUCCESS AND FLORAL EVOLUTION IN MIMULUS GUTTATUS by Gerardo Arceo-Gómez B.S., University of Yucatan, 2005 M.S., Institute of Ecology, 2008 Submitted to the Graduate Faculty of the Kenneth P. Dietrich School of Arts and Sciences in partial fulfillment of the requirements for the degree of Doctor of Philosophy, Biological Sciences University of Pittsburgi h 2014 UNIVERSITY OF PITTSBURGH KENNETH P. DIETRICH SCHOOL OF ARTS AND SCIENCES This dissertation was presented by Gerardo Arceo-Gómez It was defended on February 26, 2014 and approved by Dr. Susan Kalisz, Dept. of Biological Sciences, University of Pittsburgh Dr. Randall J. Mitchell, Dept. of Biology, University of Akron Dr. Brian Traw, Dept. of Biological Sciences, University of Pittsburgh Dr. Stephen Tonsor, Dept. of Biological Sciences, University of Pittsburgh Dissertation Advisor: Dr. Tia-Lynn Ashman, Dept. of Biological Sciences, University of Pittsburgh ii Copyright © by Gerardo Arceo-Gómez 2014 iii UNCOVERING THE CONSEQUENCES OF CO-FLOWERING AND POLLINATOR SHARING: EFFECTS OF LOCAL COMMUNITY CONTEXT ON POLLEN TRANSFER DYNAMICS, FEMALE REPRODUCTIVE SUCCESS AND FLORAL EVOLUTION IN MIMULUS GUTTATUS Gerardo Arceo-Gómez, PhD University of Pittsburgh, 2014 While plant-pollinator interactions commonly take place within a larger community context, studies of plants and their pollinators have typically focused on pair-wise interactions. Co- flowering species whithin multi-species communities may influence pollinator foraging decisions and hence plant reproductive success and floral evolution. For instance, plants growing in highly diverse areas may be more pollen limited than plants in species-poor areas due to the high levels of pollinator competition and interspecific pollen transfer. As a result, stronger selection pressures can also be expected in highly diverse areas in order to increase the quantity and/or quality of pollen reaching conspecific stigmas. However, how plant community composition contributes to the severity of pollen limitation, what the potential underlying mechanisms are and how selection on floral traits changes with increasing community diversity is still unclear. In this study I use Mimulus guttatus as a model system to evaluate the effect of local co-flowering community context on quantity and quality aspects of pollen limitation, pollen transfer dynamics, heterospecific pollen effects and selection processes in high and low diversity areas. I show that the relative contribution of pollen quantity and quality limitation to overall pollen limitation of reproductive success depends on the co-flowering community context in which M. guttatus exists. I further uncover heterospecific pollen receipt as a potential mechanism underlying decreased reproductive success in highly diverse areas by showing that complex iv interactions among multiple heterospecific pollen donors can exacerbate its effects and that heterospecific pollen receipt can have an even greater detrimental effect on self compared to outcross conspecific pollen. Finally, I show that co-flowering community context can be an important driver of selection that promotes floral trait differentiation among populations, in the case of M. guttatus in flower longevity. By combining observational, experimental, field and greenhouse approaches, this study extends our knowledge of the processes underlying insufficient pollination in natural communities, reveals new complexities in our understanding of heterospecific pollen effects and advances our understanding of the community properties shaping the evolutionary dynamics of constituent populations. v TABLE OF CONTENTS INTRODUCTION......................................................................................................................... 1 1.0 PATTERNS OF POLLEN QUANTITY AND QUALITY LIMITATION OF PRE-ZYGOTIC REPRODUCTION IN MIMULUS GUTTATUS VARY WITH CO- FLOWERING COMMUNITY CONTEXT ............................................................................... 5 1.1 INTRODUCTION ............................................................................................... 5 1.2 MATERIALS AND METHODS ...................................................................... 10 1.2.1 Study system ................................................................................................... 10 1.2.2 Patterns of pollen quantity and quality limitation in high- and low- diversity seeps ............................................................................................................. 11 1.3 RESULTS ........................................................................................................... 15 1.4 DISCUSSION ..................................................................................................... 16 1.5 CONCLUSIONS ................................................................................................ 20 2.0 HETEROSPECIFIC POLLEN DEPOSITION: DOES DIVERSITY ALTER THE CONSEQUENCES? .......................................................................................................... 23 2.1 INTRODUCTION ............................................................................................. 23 2.2 MATERIALS AND METHODS ...................................................................... 27 2.2.1 Study system ................................................................................................... 27 vi 2.2.2 Experimental plant material ......................................................................... 28 2.2.3 Pollination treatments ................................................................................... 29 2.2.4 Components of female reproductive success ............................................... 30 2.2.5 Data analyses .................................................................................................. 31 2.3 RESULTS ........................................................................................................... 32 2.3.1 Overall pollen load composition and conspecific load size effects ............ 32 2.3.2 Diversity and identity effects on stigma reopening and fertile seed production ................................................................................................................... 33 2.3.3 Multi-species interactions on fertile seed production ................................. 33 2.3.4 Mechanisms: seed abortion and pollen tube growth interference ............ 34 2.4 DISCUSSION ..................................................................................................... 35 2.4.1 What are the effects of diversity in the heterospecific load? ..................... 35 2.4.2 What is the role of CP loss in M. guttatus reproductive success? .............. 36 2.4.3 What are the mechanisms involved in fertilization failure? ...................... 36 2.4.4 Ecological and evolutionary implications in natural communities ........... 37 3.0 CO-FLOWERING COMMUNITY CONTEXT INFLUENCES FEMALE FITNESS AND ALTERS THE ADAPTIVE VALUE OF FLOWER LONGEVITY IN MIMULUS GUTTATUS ............................................................................................................. 43 3.1 INTRODUCTION ............................................................................................. 43 3.2 MATHERIALS AND METHODS ................................................................... 48 3.2.1 Co-flowering community diversity ............................................................... 48 3.2.2 Pollinator visitation ....................................................................................... 49 3.2.3 Pollen receipt .................................................................................................. 50 vii 3.2.4 Co-variation between floral traits and community context ....................... 51 3.2.5 Adaptive value of flower longevity ............................................................... 51 3.2.6 Pre-planned contrasts.................................................................................... 53 3.3 RESULTS ........................................................................................................... 55 3.3.1 Co-flowering community diversity ............................................................... 55 3.3.2 Pollinator visitation ....................................................................................... 55 3.3.3 Pollen receipt .................................................................................................. 56 3.3.4 Co-variation between floral traits and community context ....................... 56 3.3.5 Adaptive value of flower longevity ............................................................... 57 3.4 DISCUSSION ..................................................................................................... 58 3.4.1 Co-flowering diversity effects on the pollination environment ................. 59 3.4.2 Co-flowering diversity effects on the adaptive value of flower longevity . 61 3.4.3 Why flower longevity?................................................................................... 63 3.4.4 Species richness versus sampling and phylogenetic effects ....................... 65 4.0 HETEROSPECIFIC POLLEN RECEIPT AFFECTS SELF POLLEN MORE THAN OUTCROSS POLLEN IN MIMULUS GUTTATUS: IMPLICATIONS FOR MIXED-MATING PLANTS .....................................................................................................
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